基于分子标记辅助选择的玉米导入系群体构建与自交系改良
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摘要
分子标记辅助选择(molecular marker-assisted selection:MAS)是现代生物育种体系的主要领域,完善MAS的理论与技术体系对有效提高玉米遗传改良水平具有重要意义。在回交转育程序中利用MAS技术构建染色体片段代换系或染色体片段导入系群体,不仅可缩短选育年限,提高选育效果,而且可获得覆盖全基因组的、片段衔接的全套近等基因系,直接为基因定位、克隆和理论及应用研究提供基础材料。
本研究选用强优势杂交组合豫玉22的亲本87-1和Zong3为受体亲本,以糯质、抗病性和抗倒伏性较好的HB522为供体亲本,经杂交和连续3代回至BC3F1后,自交得到BC3F2。从BC1代开始,每个回交世代利用导入片段两侧分子标记进行导入片段的前景选择,利用覆盖全基因组SSR标记进行轮回亲本的背景选择,构建单片段导入系基础群体,并对其遗传背景、导入片段大小、数目和覆盖率及其表型效应等进行评价,对几个重要的产量性状QTL进行初步定位,同时,通过三种选择策略得到了waxy基因和Rf3基因的辅助回交改良系,并对改良系的前景选择效果、背景效应及配合力效应进行评价。取得的初步结果如下：
1、在87-1×HB522的BC3F1群体中筛选到83个遗传背景回复率高的单株,自交构建成含有40个目标片段的单片段导入系基础群体87-1HBSSILs。在综3×HB522的BC3F1群体中筛选到122个高遗传背景回复率的个体,自交构建成含有78个目标片段的导入系群体,其后经过进一步回交、自交,直到BC4F3,最终构建了98个Z3HBSSILs基础群体。
2、在87-1HBSSILs群体中,各导入系中所导入的片段数目不等,平均导入4个片段；相对于参考图谱IBM neighbor2005,导入片段长度在0.03～342.8cM之间,平均长度为91.1cM,导入片段覆盖48.96%玉米基因组。Z3HBSSILs导入系群体共导入188个供体亲本的基因组片段,不同导入系含有的供体片段数目1-6个不等,平均每个导入系有导入片段1.92个；相对于参考图谱IBM neighbor2005,导入片段的长度在4.8cM-256cM,平均长度为63.7cM,导入片段共覆盖86.2%的玉米基因组。
3、在保定和张家口两地,对Z3HBSSILs群体的穗行数、行粒数、百粒重、穗粒重进行鉴定,基于导入系群体的表型,开展QTL鉴定。两个环境下共检测到15个穗行数QTL,单个QTL可解释4.57%～33.06%的表型变异,其中2个QTL在两地均被检测到；6个行粒数QTL,单个QTL可解释的表型变异为4.57%～9.86%；9个百粒重QTL,单个QTL可解释的表型变异为4.24%～19.27%,其中,1个QTL在两地均被检测；以及2个穗粒重QTL,分别解释6.493%和10.131%的表型变异,2个QTL在两个环境均被检测到。
4、基于三种分子标记辅助选择(MAS)策略,针对Rf3和waxy(wx)基因开展了基于MAS的回交转育,获得了3个改良系。分别对这3个改良系目标性状的改良效果进行分析证实,Rf3基因的导入可使S-Mo17(rf3rf3)育性恢复；导入waxy基因,不仅没有改变总淀粉含量,而且显著提高了支链淀粉含量,降低了直链淀粉含量。
5、对三种MAS选择策略得到的3个改良系的背景评价发现,(1)回交各世代进行标记背景选择的改良系S-Mo17(Ry3Rf3),背景回复率为96.0%,改良系中供体染色体片段残留仅为4%,分布于第7、8和9染色体的个别区域；(2)在BC3F1进行一次标记辅助的背景选择,所得改良系Z3-wxwx的背景回复率为94.6%,改良系中存在5.4%的遗传累赘和残留,分布在目标基因wx所在的第9染色体以及第3、6、7染色体上；(3)在BC3F1只进行表型选择而不进行标记辅助的背景选择,所得改良系87-1-wxwx的背景回复率仅为82.1%,改良系中存在17.9%的供体残留,除第1、10染色体外,其他8条染色体上均有背景残留。从表型来看,S-Mo17(Rf3Ry3)比S-Mo17(rf3rf3)株高变矮6.0cm,开花期提早3天；87-1-wwx比87-1株高和穗位高分别降低23cm和6.7cm、穗长变短,抽雄期提早3天；Z3-wxwx比Z3株高增加19.3cm、穗位高增加6.7cm,开花期推迟3天,87-1-wwx与87-1间差异较显著。由此表明,背景的回复率越低,改良系与原自交系的性状差异就越大,与表型选择相比,通过SSR标记进行背景选择可有效降低供体非目标片段的导入。
6、改良前后自交系与多个玉米自交系杂交,组配杂交组合,以检测供体染色体片段残留对自交系的配合力和杂交种性状的影响,结果发现,S-Mo17(rf3rf3)和S-Mo17(Rf3Rf3)、Z3和Z3-wxwx、87-1和87-1-wxwx分别与5个测验种杂交组合的表现型相似,改良前后的自交系表现出相似的一般配合力和特殊配合力,进一步地说明利用回交结合分子标记辅助选择,既可有效地改良目标性状,又不至于改变优良自交系的配合力特性。但由于残留遗传背景的干扰,改良前后自交系与测验种所组配的相似组合间,仍然存在个别性状的差异。
The molecular marker-assisted selection (MAS) is an useful tool in crop breeding program and plays an important role in maize genetic improvement. The chromosome segment substitution lines (CSSLs) or chromosome segment introgression lines (CSILs) population which developed using MAS during backcross program can not only shorten the breeding period and raise the selecting efficiency, but also get the whole set of near-isogenic lines(NILs) with linked segment covering the whole genome which can be used as basic material for gene mapping, gene cloning and other theoretical and practical research.
In this study, Zong3and87-1, parents of an elite maize hybrid Yuyu22, were used as recipient line, respectively, and HB522, an elite waxy inbred with disease resistance and lodging resisitence, was selected as donor line. Two set of link-up single segment introgression line (SSIL) populations were developed through3cycle advanced backcrossing.Foreground selection were conducted using the two SSR markers defined the introgressed segment and background selection of populations were screened using genome-wide SSR markers in each cycle of backcross. The size and number of introgressed segment, coverage of genome and phenotype were evaluated. QTL mapping for yield and yield-association traits were also performed. Additionally, R/3and waxy were used to improve inbred lines using three strategies of MAS, and background in these improved lines were screeded by genome-wide SSR markers, the effects of phenotype and combining ability caused by residual segment of donor were evaluated of improved lines and their hybrids. The results were shown as followling:
1. The40lines were developed by83individuals derived from BC3F1of crossing87-1×HB522, and98lines were developed by122individuals derived from BC4F3of crossing Zong3×HB522, respectively. The population developed from87-1as recurrent parent was defined as87-1HBSSILs, similarly, the population developed from Zong3as recurrent parent was defined as Z3HBSSILs.
2. A total of40donor genomic segments were introgressed into the87-1inbred. An average length of single introgressed segment was91.10cM on the basis of reference map IBM neighbor2005, ranged from0.03to342.86cM and covered a48.96%genome of87-1. A total of188donor segments were developed by the Z3HBSSILs. The number of these introgressed segments in each introgressed line varies from1to6, while the average number of them is1.92. The average length of introgressed segments was63.7cM, ranging from4.8cM to256cM on the basis of reference map IBM neighbor2005. The introgressed segment covered86.2%genome of maize.
3. The QTL for grain yield and yield-association traits were identified using Z3HBSSIL population in Baoding and Zhangjiakou. A total of32QTL were detected across two environments, of which15for row number per ear (RNPE),6for kernel number per row (KPR),9for grain weight (GW) and2for weight per ear (WE), respectively. Each QTL explaining from4.57～33.06%,4.57～9.86%,4.24～19.27%and6.493～10.131%of phenotypic variance of RNPE, RNPE, KW and WE, respectively. The QTL around bnlg1083was identified in two traits across two locations.
4. Three improved lines were developed based on three MAS strategies for Rf3and waxy gene selecting. Improved effect were evaluated and indicated that the improved line S-Mo17Rf3Rf3was completely fertile because of the introgression of Rf3gene. The total starch content of Z3-wxwx kernel was identical to that of in the Zong3kernel, but the amylopectin content of the former was significantly increased and its amylase content dramatically decreased because of the introgression of wx gene.
5. Comparison of residual background among3improved lines derived from different MAS strategies indicated that17.9%donor chromosomal fragment was retained in87-1-wxwx which was developed by only selection of phenotype in BC3F1generation, and the residual segment distributed on all8chromosomes except chromosome1and10. While only about4%chromosomal residual was detected and distributed on minor region of chromsome7,8and9in Mo17Rf3Rf3which was developed by screening background using whole genome SSR markers in each generation. The background residual of Z3-wxwx line which was developed by background selection in BC3F1using MAS, was less than that of the line of87-1-wxwx, but more than that of Mo17Rf3Rf3. Phenotype were evaluated and revealed that the day to flowering (DTF) of S-Mo17Rf3Rf3was3days earlier than that of S-Mo17rf3rf3and its plant height (PH) was6.0cm lower than that of S Mo17rf3rf3. The PH and ear height (EH) of87-1-wxwx were both lower than that of87-1and its ear length (EL) and DTF was slightly shorter and3days earlier than that of87-1, respectively. The PH and EH of Z3-wxwx were taller than that of Z3and its DTF was3days later than that of Z3. The significant difference (P<0.05)) between87-1and87-1-wxwx was identified based on performance of multiple traits. Taken together, these results indicated that background selection by MAS could reduce the residual of non-target fragment of donor.
6. Combining ability performed sililar between the MAS-derived lines and their corresponding recurrent lines. It was indicated that combining MAS and backcross could both improve the target traits and keep the combining ability of inbred line. However, there were still differences in few traits between the combinations of improved and unimproved inbred line with testers due to the interference of the residual background genome from the donor.
本研究选用强优势杂交组合豫玉22的亲本87-1和Zong3为受体亲本,以糯质、抗病性和抗倒伏性较好的HB522为供体亲本,经杂交和连续3代回至BC3F1后,自交得到BC3F2。从BC1代开始,每个回交世代利用导入片段两侧分子标记进行导入片段的前景选择,利用覆盖全基因组SSR标记进行轮回亲本的背景选择,构建单片段导入系基础群体,并对其遗传背景、导入片段大小、数目和覆盖率及其表型效应等进行评价,对几个重要的产量性状QTL进行初步定位,同时,通过三种选择策略得到了waxy基因和Rf3基因的辅助回交改良系,并对改良系的前景选择效果、背景效应及配合力效应进行评价。取得的初步结果如下：
1、在87-1×HB522的BC3F1群体中筛选到83个遗传背景回复率高的单株,自交构建成含有40个目标片段的单片段导入系基础群体87-1HBSSILs。在综3×HB522的BC3F1群体中筛选到122个高遗传背景回复率的个体,自交构建成含有78个目标片段的导入系群体,其后经过进一步回交、自交,直到BC4F3,最终构建了98个Z3HBSSILs基础群体。
2、在87-1HBSSILs群体中,各导入系中所导入的片段数目不等,平均导入4个片段；相对于参考图谱IBM neighbor2005,导入片段长度在0.03～342.8cM之间,平均长度为91.1cM,导入片段覆盖48.96%玉米基因组。Z3HBSSILs导入系群体共导入188个供体亲本的基因组片段,不同导入系含有的供体片段数目1-6个不等,平均每个导入系有导入片段1.92个；相对于参考图谱IBM neighbor2005,导入片段的长度在4.8cM-256cM,平均长度为63.7cM,导入片段共覆盖86.2%的玉米基因组。
3、在保定和张家口两地,对Z3HBSSILs群体的穗行数、行粒数、百粒重、穗粒重进行鉴定,基于导入系群体的表型,开展QTL鉴定。两个环境下共检测到15个穗行数QTL,单个QTL可解释4.57%～33.06%的表型变异,其中2个QTL在两地均被检测到；6个行粒数QTL,单个QTL可解释的表型变异为4.57%～9.86%；9个百粒重QTL,单个QTL可解释的表型变异为4.24%～19.27%,其中,1个QTL在两地均被检测；以及2个穗粒重QTL,分别解释6.493%和10.131%的表型变异,2个QTL在两个环境均被检测到。
4、基于三种分子标记辅助选择(MAS)策略,针对Rf3和waxy(wx)基因开展了基于MAS的回交转育,获得了3个改良系。分别对这3个改良系目标性状的改良效果进行分析证实,Rf3基因的导入可使S-Mo17(rf3rf3)育性恢复；导入waxy基因,不仅没有改变总淀粉含量,而且显著提高了支链淀粉含量,降低了直链淀粉含量。
5、对三种MAS选择策略得到的3个改良系的背景评价发现,(1)回交各世代进行标记背景选择的改良系S-Mo17(Ry3Rf3),背景回复率为96.0%,改良系中供体染色体片段残留仅为4%,分布于第7、8和9染色体的个别区域；(2)在BC3F1进行一次标记辅助的背景选择,所得改良系Z3-wxwx的背景回复率为94.6%,改良系中存在5.4%的遗传累赘和残留,分布在目标基因wx所在的第9染色体以及第3、6、7染色体上；(3)在BC3F1只进行表型选择而不进行标记辅助的背景选择,所得改良系87-1-wxwx的背景回复率仅为82.1%,改良系中存在17.9%的供体残留,除第1、10染色体外,其他8条染色体上均有背景残留。从表型来看,S-Mo17(Rf3Ry3)比S-Mo17(rf3rf3)株高变矮6.0cm,开花期提早3天；87-1-wwx比87-1株高和穗位高分别降低23cm和6.7cm、穗长变短,抽雄期提早3天；Z3-wxwx比Z3株高增加19.3cm、穗位高增加6.7cm,开花期推迟3天,87-1-wwx与87-1间差异较显著。由此表明,背景的回复率越低,改良系与原自交系的性状差异就越大,与表型选择相比,通过SSR标记进行背景选择可有效降低供体非目标片段的导入。
6、改良前后自交系与多个玉米自交系杂交,组配杂交组合,以检测供体染色体片段残留对自交系的配合力和杂交种性状的影响,结果发现,S-Mo17(rf3rf3)和S-Mo17(Rf3Rf3)、Z3和Z3-wxwx、87-1和87-1-wxwx分别与5个测验种杂交组合的表现型相似,改良前后的自交系表现出相似的一般配合力和特殊配合力,进一步地说明利用回交结合分子标记辅助选择,既可有效地改良目标性状,又不至于改变优良自交系的配合力特性。但由于残留遗传背景的干扰,改良前后自交系与测验种所组配的相似组合间,仍然存在个别性状的差异。
The molecular marker-assisted selection (MAS) is an useful tool in crop breeding program and plays an important role in maize genetic improvement. The chromosome segment substitution lines (CSSLs) or chromosome segment introgression lines (CSILs) population which developed using MAS during backcross program can not only shorten the breeding period and raise the selecting efficiency, but also get the whole set of near-isogenic lines(NILs) with linked segment covering the whole genome which can be used as basic material for gene mapping, gene cloning and other theoretical and practical research.
In this study, Zong3and87-1, parents of an elite maize hybrid Yuyu22, were used as recipient line, respectively, and HB522, an elite waxy inbred with disease resistance and lodging resisitence, was selected as donor line. Two set of link-up single segment introgression line (SSIL) populations were developed through3cycle advanced backcrossing.Foreground selection were conducted using the two SSR markers defined the introgressed segment and background selection of populations were screened using genome-wide SSR markers in each cycle of backcross. The size and number of introgressed segment, coverage of genome and phenotype were evaluated. QTL mapping for yield and yield-association traits were also performed. Additionally, R/3and waxy were used to improve inbred lines using three strategies of MAS, and background in these improved lines were screeded by genome-wide SSR markers, the effects of phenotype and combining ability caused by residual segment of donor were evaluated of improved lines and their hybrids. The results were shown as followling:
1. The40lines were developed by83individuals derived from BC3F1of crossing87-1×HB522, and98lines were developed by122individuals derived from BC4F3of crossing Zong3×HB522, respectively. The population developed from87-1as recurrent parent was defined as87-1HBSSILs, similarly, the population developed from Zong3as recurrent parent was defined as Z3HBSSILs.
2. A total of40donor genomic segments were introgressed into the87-1inbred. An average length of single introgressed segment was91.10cM on the basis of reference map IBM neighbor2005, ranged from0.03to342.86cM and covered a48.96%genome of87-1. A total of188donor segments were developed by the Z3HBSSILs. The number of these introgressed segments in each introgressed line varies from1to6, while the average number of them is1.92. The average length of introgressed segments was63.7cM, ranging from4.8cM to256cM on the basis of reference map IBM neighbor2005. The introgressed segment covered86.2%genome of maize.
3. The QTL for grain yield and yield-association traits were identified using Z3HBSSIL population in Baoding and Zhangjiakou. A total of32QTL were detected across two environments, of which15for row number per ear (RNPE),6for kernel number per row (KPR),9for grain weight (GW) and2for weight per ear (WE), respectively. Each QTL explaining from4.57～33.06%,4.57～9.86%,4.24～19.27%and6.493～10.131%of phenotypic variance of RNPE, RNPE, KW and WE, respectively. The QTL around bnlg1083was identified in two traits across two locations.
4. Three improved lines were developed based on three MAS strategies for Rf3and waxy gene selecting. Improved effect were evaluated and indicated that the improved line S-Mo17Rf3Rf3was completely fertile because of the introgression of Rf3gene. The total starch content of Z3-wxwx kernel was identical to that of in the Zong3kernel, but the amylopectin content of the former was significantly increased and its amylase content dramatically decreased because of the introgression of wx gene.
5. Comparison of residual background among3improved lines derived from different MAS strategies indicated that17.9%donor chromosomal fragment was retained in87-1-wxwx which was developed by only selection of phenotype in BC3F1generation, and the residual segment distributed on all8chromosomes except chromosome1and10. While only about4%chromosomal residual was detected and distributed on minor region of chromsome7,8and9in Mo17Rf3Rf3which was developed by screening background using whole genome SSR markers in each generation. The background residual of Z3-wxwx line which was developed by background selection in BC3F1using MAS, was less than that of the line of87-1-wxwx, but more than that of Mo17Rf3Rf3. Phenotype were evaluated and revealed that the day to flowering (DTF) of S-Mo17Rf3Rf3was3days earlier than that of S-Mo17rf3rf3and its plant height (PH) was6.0cm lower than that of S Mo17rf3rf3. The PH and ear height (EH) of87-1-wxwx were both lower than that of87-1and its ear length (EL) and DTF was slightly shorter and3days earlier than that of87-1, respectively. The PH and EH of Z3-wxwx were taller than that of Z3and its DTF was3days later than that of Z3. The significant difference (P<0.05)) between87-1and87-1-wxwx was identified based on performance of multiple traits. Taken together, these results indicated that background selection by MAS could reduce the residual of non-target fragment of donor.
6. Combining ability performed sililar between the MAS-derived lines and their corresponding recurrent lines. It was indicated that combining MAS and backcross could both improve the target traits and keep the combining ability of inbred line. However, there were still differences in few traits between the combinations of improved and unimproved inbred line with testers due to the interference of the residual background genome from the donor.
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